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Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction
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Cognitive Development
Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction Chris Sinha School of Foreign Languages, Hunan University and Department of Linguistics, Lund University, Sweden
a r t i c l e
i n f o
Keywords: Biocultural niche construction Epigenesis Ontogenesis Praxis Semiosphere
a b s t r a c t In this article I address the significance of semiotic processes supporting early social interaction, communication and learning in the evolution of the modern human niche of infancy and childhood, known to be extended even in comparison with closely related hominin species. Human infancy and childhood is a biocultural niche, embedded within and causally contributing to the expansion and elaboration of the wider human biocultural complex, including both semiotic and praxic spheres. Epigenetic constructive processes were crucial in the evolution of the niche of ontogenesis, and niche construction through epigenetic augmentation is the key to understanding human symbolic evolution, the advent of human behavioral modernity and the capture of evolutionary processes by socio-cultural dynamics. © 2015 Elsevier Inc. All rights reserved.
1. Introduction [It] is always difficult for the psychologist to think of anything ‘existing’ in a culture . . . We are, alas, wedded to the idea that human reality exists within the limiting boundary of the human skin! (Bruner, Olver, & Greenfield, 1966: 321) The aim of this article is to develop a hypothesis that this author first advanced, in general terms, more than a quarter century ago: “Rather than seeing cultural evolution as “taking off” from a terminal point of biological evolution, we should rather see evolutionary biological processes as having been “captured” by an emergent cultural process, with ontogenetic processes (especially those involving representation, symbolization and communication) as a crucial catalyst and product of the co-evolution of culture and biology . . . “The evolution of infancy was the biological mechanism through which the potential for intergenerational cultural transmission created by tool use was optimized” (Sinha, 1988: 104, 108). In fleshing out this hypothesis I attempt to draw together ideas and arguments from biocultural niche construction theory, research in the culturally driven evolution of human cognition and language, research on infant communication, palaeontological research on the evolution of the human life span and archaeological evidence of human symbolic evolution. The general perspective that I adopt stresses the co-articulation and dynamic co-evolution of three inter-related and distinctively human niches; namely, ontogenesis, semiosis, and praxis. In drawing these threads together, I will advance a defence of two key hypotheses advanced by Arbib (2012), namely that: (a) there is an intimate connection between the neurocognitive evolution of praxic action, and that of symbolic communication and language; and (b) that “language proper” (what I shall call evolutionary modern language) was a relatively late evolutionary accomplishment, significantly postdating the speciation of homo sapiens.
E-mail address: [email protected] http://dx.doi.org/10.1016/j.cogdev.2015.09.006 0885-2014/© 2015 Elsevier Inc. All rights reserved.
Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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2. Ontogenesis: the niche of infancy and childhood The role of social learning of abilities acquired in infancy and childhood in the evolution of modern human social and cognitive capacities has been foregrounded in much recent research (Richerson and Boyd, 2005; Van Schaik and Burkart, 2011). Social learning of complex skill repertoires, including language, is optimized by the extended childhood of our species; as Arbib (2012: 166) observes, “the prolonged period of infant dependency . . . combines with the willingness of adults to act as caregivers and the consequent development of social structures to provide the conditions for complex social learning”. The developmental life history of the homo sapiens species is well known to be significantly extended in comparison with extant ape species. The biology of ontogenetic life history is assessed in comparative research by physiological and anatomical markers. Although there is disagreement about whether the evolutionary process leading to the modern human life history involved heterochrony (Gould, 1977; Parker and McKinney, 2012) or the insertion of new biological maturational stages (Locke and Bogin, 2006), there is a consensus that its product—extended life history—did not appear in human evolution until the emergence of later members of the genus Homo (Dean, 2007; Lee, 2012; Robson and Wood, 2008; Schwartz, 2012). There is evidence that the process of extension continued through to the emergence of our own species, and (although this conclusion is not undisputed) that the early stages of modern human life are uniquely extended, even in comparison with Neanderthals. Tanya Smith, on the basis of her and her colleagues’ comparative dentition studies of early Homo sapiens sapiens and Homo sapiens neanderthalis (henceforth modern human and Neanderthal) child fossil remains (Smith et al., 2007; Smith, Toussaint, Reid, Olejniczac, & Hublin, 2007), concludes that “the characteristically prolonged development of living humans fully evolved after these taxa diverged” (Smith, 2013: 203), signalling the contemporaneous “advent of corresponding social, biological, and cultural changes necessary to support highly dependent children with prolonged opportunities for social learning in early childhood” (Smith et al., 2007: 6132). Infancy, childhood and adolescence are not merely biological stages of organismic development. They are also developmental stages of a biocultural niche supporting mutual, intersubjective, emotional, communicative and cognitive engagement (Trevarthen, this issue). In the course of human development, the proximal environment of the organism develops along with the maturational processes underlying the individual’s development. This developmental, and developing, niche includes as its most important component other human beings, including the caregiving adults and older children who participate in the co-construction of its material, communicative and symbolic properties. This co-construction process involves (a) the spatio-temporal organization of settings involving joint actions contributing to the reproduction of socio-cultural habitus (Bourdieu, 1977); and (b) the scaffolding and shaping of interactions in the niche on the part of the caregiver (Ninio and Bruner, 1978; Wood, Bruner, & Ross, 1976), in response to both the actions and the competences of the developing child. These fundamental processes in niche construction and niche development can be thought of as involving the cultural and situational structuring of affordances for learning, and are the socio-developmental mechanisms supporting inter-generational cultural transmission in all human cultures. Evolutionary and developmental processes take place on different time scales or durées, of phylogenesis, sociogenesis (or sociocultural evolution), ontogenesis and microgenesis (Valsiner and van der Veer, 2000). Although, as I stressed above, the niche develops along with the developing human being, the time scale that principally characterizes niche construction and niche development processes is not that of ontogenesis, but that of microgenesis. Microgenesis as individual or collaborative situated learning and development also implicates the time scale of actual interaction, labelled enchrony by Enfield (2011, 2013), in distinction to the traditional structural linguistic distinction between diachrony and synchrony. “An enchronic perspective on human communication focuses on sequences of interlocking or interdependent communicative moves that are taken to be co-relevant, and causally-conditionally related. Enchrony is a level of temporal-causal grain (typically, ‘conversational time’) that an analyst of communication can adopt, as distinct from other possible perspectives, fitted to other purposes, that focus on other temporal scales and other kinds of causal-conditional process; these include phylogenetic, diachronic, ontogenetic, epigenetic and synchronic perspectives” (Enfield, 2011: 287; see also Steffensen and Pedersen, 2013). Enchrony, in Enfield’s definition, is prototypically discursive temporality. From an ontogenetic perspective, however, enchronic interactional coordination is manifested long before the emergence in development of language, and is a fundamental property of infant as well as adult communicative meaning-making (Trevarthen, this issue). It can be argued, furthermore, that the notion of enchrony can equally be applied to the planning, sequencing and timing of component actions whose combination makes up complex praxic actions (Arbib, 2012), including cooperative praxis. If this is correct, enchrony is, in general, the durée that most appropriately characterizes human skilled action and interaction, and that is in at least some circumstances co-temporaneous with microgenesis, the appropriation and mastery of new actions and new communicative resources. We can visualize the different, but inter-articulated time scales as being embedded one within another, as in Fig. 1. Each level provides a context and platform for the levels that are subsequently embedded, but this dependency is not one of unidirectional determination. The processes that dynamically unfold in the different time scales are not independent of each other. Ontogenesis, for example, evolves phylogenetically and, through epigenesis and Baldwin effects (Sinha, 1988 Ch. 4), phylogenetic evolution itself is mediated by ontogenetic development. A further clarification of the relationship between microgenesis and enchrony is called for here. Microgenesis implicates and is supported by enchrony; but not all instances of enchronically governed action and interaction are simultaneously instances of microgenesis.
Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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Fig. 1. Time scales in evolution, development, learning, action and communication.
My argument, then, is that human infancy and childhood is a developmental (and developing) niche that evolved in our species as an adaptation to the enchronic properties of human action and interaction, thereby maximizing the human microgenetic learning potential for the cultural transmission of both language and skilled praxic action. To further specify this proposal, I next review recent developments in evolutionary niche construction theory.
3. Niche construction, semiosphere and technosphere Niche construction theory (Laland, Odling-Smee, & Feldman 2000; Odling-Smee, Laland, & Feldman 2003) is a relatively new approach in evolutionary biology that seeks to integrate an ecological dimension into the Darwinian theory of evolution by natural selection. This theory is regarded by many evolutionary biologists as providing a significant revision of the Neo-Darwinian “modern synthesis” that unified Darwin’s theory of natural and sexual selection with 20th century population genetics. A more radical interpretation of niche construction theory sees it as offering a non-reductionist Darwinian alternative to Neo-Darwinism. In the Neo-Darwinian synthesis, the unit of selection (what is selected) is the gene, or more specifically alternative variants (alleles) of the “same” gene. The agent of selection (what does the selecting) is the extra-organismic environment, including (a) the inanimate surround, (b) other species (a and b together being the basis of natural selection); together with (c) (subpopulations of) genes of the same species (the basis of sexual and kin selection). The relevant attribute upon which selection works (what is selected for) is any genetically transmitted trait. The mechanism of selection determines the differential reproductive success of the gene (allele) within the population of interacting genes, and thus the frequency distributions of genes and traits in the population. This model, when appropriately formalized, can be extended by including cultural traits in the environment, that act as “amplifiers” on the selection of genetic variation: this is known as the theory of gene-culture co-evolution (Lumsden and Wilson, 1981). Genes do not, of course, come singly, but as combinations (genotypes), “packaged” in organisms (phenotypes). It is this distinction that Dawkins (1976) recasts as a distinction between the “replicator” (the unit of selection that is copied) and the “vehicle” (that which embodies the genotypic collection of replicators, and interacts with the environment). However, it is organisms, not genes, which are subject to direct selection pressures acting on those traits conferring fitness. The organism level of biological organization receives scant attention in population genetics but, even granted that the gene is the unit of selection, it is the organism that must be considered as the site of selection. That is to say, it is the organism, not the gene, or even the genome, that is “where the action is” in the selection of those individuals that survive. Organisms, in most (though not all) cases, can be regarded as morphological individuals. However, the actual process of selection by an “agent” occurs in relation to the functioning, behaving organism. It was for this reason that Jean Piaget (1979) upheld the leading role of behavior in evolution. In the light of this, it may be (and frequently has been) questioned to what extent it remains legitimate to identify the “replicator” with the genetic unit of selection. Even if the DNA-based biochemical replicator is the gene, the evolutionary Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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dynamic of replication-plus-selection should, it can be argued, more profitably be identified with the entire complex of the site of selection, which is the active organism in its ecological niche. This is the essential insight of niche construction theory. Ecologists emphasize that species shape, as well as being shaped by, the niches that they occupy. In Gibson’s (1979) ecological psychology, a key role is played by affordances, properties of the ecological niche affording or supporting specific kinds of action made possible by the motor system and morphology of the animal. Such actions are both species-typical (though not necessarily species unique) and adaptive. Because affordances, Gibson maintained, are directly perceived, the phenomenal world of the animal is intrinsically meaningful, in that it potentiates the activation of perception-action circuits: objects present themselves as edible, climb-able, graspable and so forth. Gibson neglected, however, to note the crucial importance of the fact that some affordances are constructed by the animal itself. In an important, and more complex, subset of such cases, the resulting niche can be seen not merely as a contingent consequence of behavior, but as a kind of quasi-artifact, inasmuch as phenotypic individuals are genetically, morphologically and behaviorally adapted to the construction of specific niches which are integral to the survival and/or reproduction strategy of the species. The term “quasi-artifact” is used here to indicate that, unlike the canonical case of human artifacts, such animal quasi-artifacts need not be produced intentionally in a process guided by representation and conceptualization. Examples of such quasi-artifactual niches are the nests of bower birds, and the dams of beavers. The male bower bird builds and decorates an elaborate nest (bower) to attract females, using attractive objects such as flowers, shells and leaves. The bower forms an integral part of the male’s mating display, and sexual selection by the female is based upon the aesthetic qualities of the bower, as much as upon the behavioral display of the male. Beavers construct, through coordinated and collaborative behavior, dams that serve both as a defence against predators, and as a means to enhance the availability of food. The dams of beavers not only serve as a constructed, quasi-artifactual niche for beavers themselves, but also reproduce the wetland ecology in which many other species thrive. As a final example of the significance of animal quasi-artifacts, we can mention the termite mound, whose material structure is not only integral to the reproductive strategy of this species of social insect, but also constitutes the morphological structure of the colony as a “group organism”. What are the implications of this integration of ecology into evolution for the Neo-Darwinian synthesis? A conservative interpretation would be that the only modification required is that the phenotype, or “vehicle”, be extended to incorporate the quasi-artifactual niche. This is, indeed, the interpretation favored by Dawkins (1982), who employs the terminology of “extended phenotype.” Under this interpretation, the “replicator” remains the gene, and only the gene. In fact, however, it is not only the gene that is copied or replicated. The niche, too, is both reproduced across generations, and serves as a fundamental precondition for genetic replication. The quasi-artifactual niche is thus both a consequence of and an agent in natural and/or sexual selection, and must then be seen as a key ingredient of the evolution of the species-typical genotype. The artifact/niche is not merely part of what is reproduced, but is also fundamental to the process of its reproduction and transmission, since it constitutes a self-made environment for adaptive selection. This co-evolutionary dynamic of reciprocal construction is captured by Laland et al., term “phenogenotypic replication”. It seems, therefore, that the integration of ecological considerations into evolutionary theory, and specifically the existence of animal quasi-artifactual niches, undermines the hard and fast distinction in the modern synthesis between germ-line and soma, genotype and phenotype, “replicator” and “vehicle”. In fact it makes better sense to say that, even granted that the unit of Darwinian selection remains the gene (allele), the “replicator” includes both the artifactual niche, and the nicheadaptive behavioral repertoire of the animal. Such considerations lead us back to Piaget’s more general proposition that behavior is the leading edge and motor of evolution, prompting the conclusion (anticipated above) that the identification by Dawkins of the “replicator” with the unit of selection (the gene, or its hypothesized cultural analogue, the “meme”) is deeply flawed, and that replication can just as well, or even better, be considered as a property of the entire site of selection—including the animal’s behavioral repertoire. In the cases discussed above, the behavioral repertoire of the species includes behaviors that are specifically adapted to the making of the quasi-artifactual niche, and these behaviors in turn support wider repertoires of behavioral strategies exploiting the niche. Quasi-artifactual niches are adaptive precisely because of the behaviors and strategies that they afford—nests are for nesting, and burrows are for burrowing. The quasi-artifactual niche in many cases can be regarded as an extension either of a behavioral repertoire (e.g., male mating display) or of the organism’s morphology (e.g., the bower bird’s bower as functionally equivalent, as an indicator of fitness, to the tail of the peacock). Indeed, we can further ask if it might be fruitful to consider certain species-specific behavioral repertoires, such as birdsong, also to be kinds of animal quasi-artifacts, inasmuch the song of the adults provides a niche within which the singing behavior is epigenetically learned (Marler and Peters, 1982; Sinha, 2004). In general terms, culturally transmitted, specialized behavioral repertoires are not merely biological, but biocultural niches, functionally equivalent to, and constructively integrated with the material dimensions of animal quasi-artifacts. If this argument is accepted, it follows that human natural languages can also be viewed as species-specific, quasi-artifactual, biocultural niches. Pinker (1994), in keeping with his nativist modularist view of the language capacity, emphatically denies that language is an artifact: he regards language as a part of the natural world, and the capacity for language as a part of human nature. We can counter Pinker’s view, however, not by insisting that languages are cultural artifacts tout court, but by regarding language as a biocultural niche and ground for what is unique about human symbolic culture. This conclusion extends the proposals advanced by Odling-Smee and Laland (2009) and Whiten and Erdal (2012), that hominid niche construction created conditions favorable for the emergence and evolution of language, by conceptualizing language itself as a constitutive (indeed the most distinctive) constituent of the evolving human biocultural complex. In other words, the human language capacity, Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. 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as a psychobiological and organismic property, did not merely evolve within a generalized human socio-cognitive niche, but can be seen as having co-evolved with the linguistic-symbolic niche created by human communication and human praxic action. As I shall argue, this apparently innocuous extension has far-reaching implications for our understanding of the ontology of language and the human language capacity. Language is a species-specific biocultural niche (Sinha, 2009; on which this section is in part based), and the ability to use and acquire it involves not only the replication of phylogenetic adaptation to this niche, but the evolution and replication of an entire symbolically mediated biocultural complex, or what Lotman (1990) called the semiosphere: the ensemble of semiotic resources implicated in the establishment, maintenance and transformation of socio-cultural habitus (Bourdieu 1977). My proposal, in a nutshell, then, is that although other species than humans may properly be said to display behaviors that can be regarded as both cultural and culturally transmitted (Whiten et al., 1999), human culture is distinguished by the predominant place occupied in it by language as a biocultural niche. Language therefore has a dual ontology (Sinha, 2006), as part of biological human species-being, “what it means to be human”, and as the foundational social institution in the Durkheimian sense (Durkheim, 1895). Furthermore, the possession of language imparts this very same dual ontology to our symbolic species (Deacon, 1997; Smolka et al., 1997); perhaps contributing, too, to a distinctively human, semiotic mode of consciousness (Arbib, 2014). Treating language as a biocultural niche yields a new perspective both on the human language capacity and on the evolution of this capacity. First, it unifies, in a non-reductionist fashion, the evolutionary dynamics of human material culture and symbolic culture. As Boivin (2008: 190) has pointed out “Tools, technologies, and other aspects of the material world of humans and their predecessors have largely been seen as the outcome of evolutionary developments, and little attempt has been made to investigate their potential role as selection forces during the course of human evolution” (my emphasis). The same can be said of the biocultural niche of language, which is not separate from the other material and symbolic components or niche-structures that make up the human biocultural complex. The biocultural niche of language is culturally situated, that is, it is dynamically embedded within the entire biocultural complex that includes both symbolic and non-symbolic artifacts. The self-constructed human biocultural complex both favored the emergence and elaboration of language, as proposed by Odling-Smee and Laland (2009: 120); and, because language is co-constitutive of the complex, was fundamentally transformed into a symbolic niche or semiosphere continuous with what we might call the materialartefactual technosphere. We can hypothesize that this process began with proto-language, almost certainly well before the emergence of anatomically and genetically modern humans some 200,000 years ago; and probably in the course of the evolution of Homo erectus from no earlier than 1.8 million years and no later than 500,000 years ago. Recent archaeological discoveries show conclusively that H. erectus individuals had both the motor manipulative skills and the semiotic capacity and motivation to engrave shells at around half a million years ago (Joordens et al., 2014). An obvious question in this context is whether such artifacts can be treated as a proxy for the emergence of evolutionary modern languages; I return to this question in the concluding section. Second, treating language as a biocultural niche means that ecological and epigenetic theories of language learning do not require the organism, as do generative linguistic accounts, to possess an innate Universal Grammar to account for language acquisition. The grammar of the language is not pre-inscribed in a language faculty; rather, grammar is the set of normative conventions regulating action within the semiosphere. This account of language and language learning is thus compatible with usage-based, cognitive-functional theories of language and language acquisition (Tomasello, 1998, 2003). The capacity for constructing a language is, from this perspective, a developing cognitive-behavioral relationship between language user and the constituents of language, just as the capacity for building a nest is a cognitive-behavioral relationship between the builder and the constituents of the nest; and it is this relationship that, in each case, has been selected for in evolution. Human natural languages are dramatically different from the communicative signals and signal systems that are ubiquitous in the animal world. The origin of this difference resides in the fact that only human languages systematically employ symbols rather than signals (Sinha, 2004). The human symbolic capacity emphatically does not take the form of a single design blueprint for grammar. Rather, its productivity and flexibility has eventuated in massive diversity of human languages and human cultures. Rejection of the hypothetical construct of Universal Grammar is consistent with recent comparative linguistic research that highlights dimensions of language variation that have in the past received insufficient attention, and which cast doubt on whether there are any true language universals at all (Evans and Levinson, 2009). Common properties of languages take the form of constraints on variation rather than putative universals, and can be hypothesized to be accounted for by common properties of the human cognitive system, in concert with the transculturally shared communicative intentions of language users. It is true that the capacity to acquire language is a transcultural and species-specific human cognitive capacity, but as I argue below this is an evolutionary outcome consequent on epigenetic plasticity in the extended life course of human infant and child development. 4. Epigenesis, enchrony and evolution Epigenesis and epigenetics are terms referring to inheritance processes and mechanisms, at different levels ranging from the molecular to the organismic, that are controlled or modulated by factors other than those inscribed in the genome (Jablonka and Lamb, 2005). Epigenesis occupies a central position in Piaget’s later work, which was strongly influenced by the theoretical biology of C.H. Waddington (e.g., Waddington, 1953). Piaget considered epigenesis to provide a third way Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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between nativism and environmentalism, and to underpin his constructivist genetic epistemology. Sinha (1988) argued that epigenetic processes provide an integrative bridge between biological and social processes in evolution and development, introducing the notion of epigenetic socio-naturalism. In this section, I outline the integrative relations between niche construction and epigenetic processes in human ontogenesis and its evolution, proposing that this dynamic coupling grounded the emergence of symbolic behaviors. Epigenetic developmental processes in ontogenetic behavioral development are those in which the developmental trajectory and final form of the developing behavior are a consequence as much of the environmental information as of the genetically encoded information. A genetically specified initial behavioral repertoire is subsequently elaborated through experience of a relevant environment, yielding an envelope of potential trajectories and outcomes. The process of elaboration is directional, and once it has taken place the initial plasticity of the embryonic, or unelaborated, repertoire is largely (though not necessarily wholly) lost. In other words, epigenesis involves a developmental transition from relative organismic plasticity and informational openness, to relative rigidity and informational closure. Augmented epigenesis is therefore, advantageous for organisms in which phenogenotypic replication involves a high level of diversity of organism-niche coupling and developmental trajectories. This is particularly relevant in the light of the diversity stressed above in human languages, and the ensuing diversity of human symbolically and linguistically mediated cultures. Regulatory genes augmenting epigenetic openness can therefore be expected to have been phenogenotypically selected for in the evolving human genome, permitting further adaptive selection for domain-specific learning in the biocultural complex, in particular for language learning and for the mimetic learning by observation of both praxic and ritual action (Donald, 1991). The class of organisms with the language capacity (normally developing human beings) can thus be theorized as a phenogenotypic replicator, systemically associated with a wider biocultural complex of interpenetrating semiosphere and technosphere, supported by symbolic and praxic-constructive cognitive capacities. Individual language acquisition and use is situated in the contexts of actuation of these inter-related capacities. What makes human symbolic culture unique is not an innate language acquisition device plus a variety of other species-specific innate cognitive modules, but a generalized semiotic capacity, epigenetically developed from a suite of cognitive capacities largely shared with other species, but attaining higher levels of organization in humans (Deacon, 1997; Piaget, 1945; Sinha, 1988, 2004). This capacity is not inscribed in the human genome, but distributed across the practices and systems co-constituting (with the epigenetically developed human organism) the human phenogenotype. The most fundamental property of these distributed and interlocking systems is, I suggest, enchrony, which is common to practices in both semiosphere and technosphere, and to those frequent cognitive-cultural practices that weave these two spheres together. As yet, we know too little to accept or reject hypotheses regarding the innateness of a specifically syntactic component of the human language capacity. We certainly should not exclude the possibility that the epigenetic processes selected in the evolution of the human biocultural complex include a predisposition for learning syntax; but this does not imply that any such predisposition is or was dedicated from the start exclusively to language. Furthermore, innate adaptations of the perceptionaction system (including the Mirror System: Arbib, 2012) facilitating the epigenetic learning of languages do not imply the innate specification of putative language universals. In an epigenetic perspective, any adaptive developmental predisposition for learning language needs neither to involve direct coding of, nor to be dedicated exclusively to, linguistic structure (Mueller, 1996). Rather, we may hypothesize that epigenetically governed adaptations initially evolved in response to proto-linguistic socio-communicative and symbolic processes, later capturing and re-canalizing behavioral adaptations (such as serial and hierarchical constructive praxis and collaborative joint action) that were initially targeted to other developmental and cognitive domains. The common factor binding these domains together is their enchronic nature, whether involving inter-individual or intraindividual temporal organization and coordination. In the materially productive realm of the technosphere, this involves the evolution of skilled praxic action, and of its learning. In the intersubjective realm of the semiosphere, this involves symbolically mediated skilled symbolic action and interaction (Fusaroli, Gangopadhyay, & Tylén, 2014), and its learning. The learning of these skilled practices is not to be understood as the processing and internalization of an “input” divorced from the learner’s activity. Rather, learning is based in the infant and child’s participation (Goodwin and Goodwin, 2004) in socially organized, enchronically structured practices, supported and constrained by the material and symbolic properties of the co-developing proximal niche. It is to the challenges posed by this socio-culturally organized, diverse, culture- and language-specific developmental process that augmented epigenesis provided the evolutionary answer. 5. Conclusion: a question of timing I conclude by suggesting that augmented epigenesis in modern human ontogenesis was not a necessary condition for the emergence of symbolic capacities and their cognitive prerequisites, which can be traced back at least half a million years. Rather, augmented epigenesis in the time-extended human ontogenetic niche furnished a phenogenotypic mechanism suitable for stabilizing and expanding the diversification and innovation in material and symbolic cultures that first appears in the archaeological record in Southern Africa in the Middle Stone Age, developing between 100,000 and 75,000 years ago (Henshilwood and Dubreuil, 2011; Henshilwood et al., 2011). This diversification and innovation is taken by many archaeologists to be the hallmark of “behavioral modernity”. Criterial for behavioral modernity is the existence of a symbolically mediated culture, in which “individuals understand that artifacts are imbued with meaning and that these meanings are Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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construed and depend on collectively shared beliefs”; which in turn “explains how human norms and conventions differ from the ritualized behaviors found in non-human primates” (Henshilwood and Dubreuil, 2011: 368–369). As noted above, there is evidence of the engraving by late H. erectus of quasi-geometric designs on shells, around half a million years ago (Joordens et al., 2014). This does not, in itself, warrant the attribution of a full-blown repertoire of behavioral modernity to these hominins. Indeed, such findings highlight the difficulties, both methodological and theoretical, that beset attributions of symbolic mediation of collectively shared beliefs to the cultures of the makers of archaeological artifacts. Nonetheless, these findings compel the conclusion that the time depth of the co-evolution of semiosphere and technosphere is far greater than that of our species. Do they also compel the conclusion that evolutionarily modern, complex language might be equally old? Just such a hypothesis was advanced by Dediu and Levinson (2013), who place the dawn of language at around half a million years ago, with the common ancestor of modern humans and Neanderthals, and suggest that presentday languages have traces of the admixture of Neanderthal languages, just as the genomes of some present-day human populations exhibit traces of interbreeding with Neanderthals1 . There is indeed evidence that Neanderthals manifested some of the practices clustered together as behavioral modernity, including personal ornamentation and funerary practices (d’Errico et al., 2012). However, there is at present no evidence that Neanderthals maintained material-symbolic “techno-complexes”, of the kind characterizing modern human populations in Southern Africa, over extended periods. It is also noteworthy that innovations introduced in the Middle Stone Age, some 80,000 years ago, by Southern African modern humans disappear from the archaeological record at about 60,000 years ago, only to be superseded by new innovations appearing some 15–20,000 years later. Some of these later innovations (from about 44,000 years ago) manifest apparent continuity with historic hunter-gatherer societies (d’Errico et al., 2012). It seems, then, that the human cultural “ratchet effect” (Tomasello, 1999), by means of which cultural innovations are preserved, transmitted and serve as the foundation for subsequent innovation, may not have been fully in place until quite late in the evolution of our species. The earliest evidence of an extended human childhood with a life history profile similar to that of contemporary children, based on analysis of the teeth of an early Homo sapiens child, is from 160,000 years ago (Smith et al., 2007). If we accept with Smith, Toussaint, Reid, Olejniczac, and Hublin (2007) that this life course was not shared by Neanderthals, it is a plausible inference that the augmented epigenetic developmental plasticity of our species was also not shared in the same degree by any other (extinct) hominin species, but evolved either as part of, or subsequent to, modern human speciation. What specific advantage, then, might species-specific augmented epigenesis have conferred, and with what consequences? Perhaps the decisive selective advantage conferred by augmented epigenesis had not only to do with the faithful copying and transmission of specific material and symbolic practices, important as this may be; but also with the role of learning and teaching in generating and tracking cycles of expansion and stabilization of the symbolic space of social and cultural structure, brought into being by the evolution of the all-pervasive, linguistically grounded semiosphere. We can think, here, perhaps of the development of socio-cognitive logics such as kinship systems and elaborate cosmologies. All of these involve a level of linguistic complexity, involving narrative (Bruner, 1990) and flexible cognitive construal (Langacker, 1987), only to be found in fully evolutionarily modern, speech-based and highly grammaticalized languages. I suggest, in conclusion, that not only did the emergence of the Homo sapiens species deliver a “language-ready brain” (Arbib, 2012), but that this brain was itself the product of a “language-ready niche of development”, whose evolution led to the marrying of augmented epigenesis with the linguistically mediated teaching and learning of social symbolic knowledge at some point in the last 80,000 years. References Arbib, M. (2012). How the brain got language: the mirror system hypothesis. Oxford: Oxford University Press. Arbib, M. (2014). Co-evolution of human consciousness and language (revisited). J. Integr. 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1 The shell engravings described by Joordens et al. were found in Java; whereas Dediu and Levinson theorize about the language capacities of an African common ancestor of modern humans and Neanderthals.
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Contents lists available at ScienceDirect
Cognitive Development
Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction Chris Sinha School of Foreign Languages, Hunan University and Department of Linguistics, Lund University, Sweden
a r t i c l e
i n f o
Keywords: Biocultural niche construction Epigenesis Ontogenesis Praxis Semiosphere
a b s t r a c t In this article I address the significance of semiotic processes supporting early social interaction, communication and learning in the evolution of the modern human niche of infancy and childhood, known to be extended even in comparison with closely related hominin species. Human infancy and childhood is a biocultural niche, embedded within and causally contributing to the expansion and elaboration of the wider human biocultural complex, including both semiotic and praxic spheres. Epigenetic constructive processes were crucial in the evolution of the niche of ontogenesis, and niche construction through epigenetic augmentation is the key to understanding human symbolic evolution, the advent of human behavioral modernity and the capture of evolutionary processes by socio-cultural dynamics. © 2015 Elsevier Inc. All rights reserved.
1. Introduction [It] is always difficult for the psychologist to think of anything ‘existing’ in a culture . . . We are, alas, wedded to the idea that human reality exists within the limiting boundary of the human skin! (Bruner, Olver, & Greenfield, 1966: 321) The aim of this article is to develop a hypothesis that this author first advanced, in general terms, more than a quarter century ago: “Rather than seeing cultural evolution as “taking off” from a terminal point of biological evolution, we should rather see evolutionary biological processes as having been “captured” by an emergent cultural process, with ontogenetic processes (especially those involving representation, symbolization and communication) as a crucial catalyst and product of the co-evolution of culture and biology . . . “The evolution of infancy was the biological mechanism through which the potential for intergenerational cultural transmission created by tool use was optimized” (Sinha, 1988: 104, 108). In fleshing out this hypothesis I attempt to draw together ideas and arguments from biocultural niche construction theory, research in the culturally driven evolution of human cognition and language, research on infant communication, palaeontological research on the evolution of the human life span and archaeological evidence of human symbolic evolution. The general perspective that I adopt stresses the co-articulation and dynamic co-evolution of three inter-related and distinctively human niches; namely, ontogenesis, semiosis, and praxis. In drawing these threads together, I will advance a defence of two key hypotheses advanced by Arbib (2012), namely that: (a) there is an intimate connection between the neurocognitive evolution of praxic action, and that of symbolic communication and language; and (b) that “language proper” (what I shall call evolutionary modern language) was a relatively late evolutionary accomplishment, significantly postdating the speciation of homo sapiens.
E-mail address: [email protected] http://dx.doi.org/10.1016/j.cogdev.2015.09.006 0885-2014/© 2015 Elsevier Inc. All rights reserved.
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2. Ontogenesis: the niche of infancy and childhood The role of social learning of abilities acquired in infancy and childhood in the evolution of modern human social and cognitive capacities has been foregrounded in much recent research (Richerson and Boyd, 2005; Van Schaik and Burkart, 2011). Social learning of complex skill repertoires, including language, is optimized by the extended childhood of our species; as Arbib (2012: 166) observes, “the prolonged period of infant dependency . . . combines with the willingness of adults to act as caregivers and the consequent development of social structures to provide the conditions for complex social learning”. The developmental life history of the homo sapiens species is well known to be significantly extended in comparison with extant ape species. The biology of ontogenetic life history is assessed in comparative research by physiological and anatomical markers. Although there is disagreement about whether the evolutionary process leading to the modern human life history involved heterochrony (Gould, 1977; Parker and McKinney, 2012) or the insertion of new biological maturational stages (Locke and Bogin, 2006), there is a consensus that its product—extended life history—did not appear in human evolution until the emergence of later members of the genus Homo (Dean, 2007; Lee, 2012; Robson and Wood, 2008; Schwartz, 2012). There is evidence that the process of extension continued through to the emergence of our own species, and (although this conclusion is not undisputed) that the early stages of modern human life are uniquely extended, even in comparison with Neanderthals. Tanya Smith, on the basis of her and her colleagues’ comparative dentition studies of early Homo sapiens sapiens and Homo sapiens neanderthalis (henceforth modern human and Neanderthal) child fossil remains (Smith et al., 2007; Smith, Toussaint, Reid, Olejniczac, & Hublin, 2007), concludes that “the characteristically prolonged development of living humans fully evolved after these taxa diverged” (Smith, 2013: 203), signalling the contemporaneous “advent of corresponding social, biological, and cultural changes necessary to support highly dependent children with prolonged opportunities for social learning in early childhood” (Smith et al., 2007: 6132). Infancy, childhood and adolescence are not merely biological stages of organismic development. They are also developmental stages of a biocultural niche supporting mutual, intersubjective, emotional, communicative and cognitive engagement (Trevarthen, this issue). In the course of human development, the proximal environment of the organism develops along with the maturational processes underlying the individual’s development. This developmental, and developing, niche includes as its most important component other human beings, including the caregiving adults and older children who participate in the co-construction of its material, communicative and symbolic properties. This co-construction process involves (a) the spatio-temporal organization of settings involving joint actions contributing to the reproduction of socio-cultural habitus (Bourdieu, 1977); and (b) the scaffolding and shaping of interactions in the niche on the part of the caregiver (Ninio and Bruner, 1978; Wood, Bruner, & Ross, 1976), in response to both the actions and the competences of the developing child. These fundamental processes in niche construction and niche development can be thought of as involving the cultural and situational structuring of affordances for learning, and are the socio-developmental mechanisms supporting inter-generational cultural transmission in all human cultures. Evolutionary and developmental processes take place on different time scales or durées, of phylogenesis, sociogenesis (or sociocultural evolution), ontogenesis and microgenesis (Valsiner and van der Veer, 2000). Although, as I stressed above, the niche develops along with the developing human being, the time scale that principally characterizes niche construction and niche development processes is not that of ontogenesis, but that of microgenesis. Microgenesis as individual or collaborative situated learning and development also implicates the time scale of actual interaction, labelled enchrony by Enfield (2011, 2013), in distinction to the traditional structural linguistic distinction between diachrony and synchrony. “An enchronic perspective on human communication focuses on sequences of interlocking or interdependent communicative moves that are taken to be co-relevant, and causally-conditionally related. Enchrony is a level of temporal-causal grain (typically, ‘conversational time’) that an analyst of communication can adopt, as distinct from other possible perspectives, fitted to other purposes, that focus on other temporal scales and other kinds of causal-conditional process; these include phylogenetic, diachronic, ontogenetic, epigenetic and synchronic perspectives” (Enfield, 2011: 287; see also Steffensen and Pedersen, 2013). Enchrony, in Enfield’s definition, is prototypically discursive temporality. From an ontogenetic perspective, however, enchronic interactional coordination is manifested long before the emergence in development of language, and is a fundamental property of infant as well as adult communicative meaning-making (Trevarthen, this issue). It can be argued, furthermore, that the notion of enchrony can equally be applied to the planning, sequencing and timing of component actions whose combination makes up complex praxic actions (Arbib, 2012), including cooperative praxis. If this is correct, enchrony is, in general, the durée that most appropriately characterizes human skilled action and interaction, and that is in at least some circumstances co-temporaneous with microgenesis, the appropriation and mastery of new actions and new communicative resources. We can visualize the different, but inter-articulated time scales as being embedded one within another, as in Fig. 1. Each level provides a context and platform for the levels that are subsequently embedded, but this dependency is not one of unidirectional determination. The processes that dynamically unfold in the different time scales are not independent of each other. Ontogenesis, for example, evolves phylogenetically and, through epigenesis and Baldwin effects (Sinha, 1988 Ch. 4), phylogenetic evolution itself is mediated by ontogenetic development. A further clarification of the relationship between microgenesis and enchrony is called for here. Microgenesis implicates and is supported by enchrony; but not all instances of enchronically governed action and interaction are simultaneously instances of microgenesis.
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Fig. 1. Time scales in evolution, development, learning, action and communication.
My argument, then, is that human infancy and childhood is a developmental (and developing) niche that evolved in our species as an adaptation to the enchronic properties of human action and interaction, thereby maximizing the human microgenetic learning potential for the cultural transmission of both language and skilled praxic action. To further specify this proposal, I next review recent developments in evolutionary niche construction theory.
3. Niche construction, semiosphere and technosphere Niche construction theory (Laland, Odling-Smee, & Feldman 2000; Odling-Smee, Laland, & Feldman 2003) is a relatively new approach in evolutionary biology that seeks to integrate an ecological dimension into the Darwinian theory of evolution by natural selection. This theory is regarded by many evolutionary biologists as providing a significant revision of the Neo-Darwinian “modern synthesis” that unified Darwin’s theory of natural and sexual selection with 20th century population genetics. A more radical interpretation of niche construction theory sees it as offering a non-reductionist Darwinian alternative to Neo-Darwinism. In the Neo-Darwinian synthesis, the unit of selection (what is selected) is the gene, or more specifically alternative variants (alleles) of the “same” gene. The agent of selection (what does the selecting) is the extra-organismic environment, including (a) the inanimate surround, (b) other species (a and b together being the basis of natural selection); together with (c) (subpopulations of) genes of the same species (the basis of sexual and kin selection). The relevant attribute upon which selection works (what is selected for) is any genetically transmitted trait. The mechanism of selection determines the differential reproductive success of the gene (allele) within the population of interacting genes, and thus the frequency distributions of genes and traits in the population. This model, when appropriately formalized, can be extended by including cultural traits in the environment, that act as “amplifiers” on the selection of genetic variation: this is known as the theory of gene-culture co-evolution (Lumsden and Wilson, 1981). Genes do not, of course, come singly, but as combinations (genotypes), “packaged” in organisms (phenotypes). It is this distinction that Dawkins (1976) recasts as a distinction between the “replicator” (the unit of selection that is copied) and the “vehicle” (that which embodies the genotypic collection of replicators, and interacts with the environment). However, it is organisms, not genes, which are subject to direct selection pressures acting on those traits conferring fitness. The organism level of biological organization receives scant attention in population genetics but, even granted that the gene is the unit of selection, it is the organism that must be considered as the site of selection. That is to say, it is the organism, not the gene, or even the genome, that is “where the action is” in the selection of those individuals that survive. Organisms, in most (though not all) cases, can be regarded as morphological individuals. However, the actual process of selection by an “agent” occurs in relation to the functioning, behaving organism. It was for this reason that Jean Piaget (1979) upheld the leading role of behavior in evolution. In the light of this, it may be (and frequently has been) questioned to what extent it remains legitimate to identify the “replicator” with the genetic unit of selection. Even if the DNA-based biochemical replicator is the gene, the evolutionary Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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dynamic of replication-plus-selection should, it can be argued, more profitably be identified with the entire complex of the site of selection, which is the active organism in its ecological niche. This is the essential insight of niche construction theory. Ecologists emphasize that species shape, as well as being shaped by, the niches that they occupy. In Gibson’s (1979) ecological psychology, a key role is played by affordances, properties of the ecological niche affording or supporting specific kinds of action made possible by the motor system and morphology of the animal. Such actions are both species-typical (though not necessarily species unique) and adaptive. Because affordances, Gibson maintained, are directly perceived, the phenomenal world of the animal is intrinsically meaningful, in that it potentiates the activation of perception-action circuits: objects present themselves as edible, climb-able, graspable and so forth. Gibson neglected, however, to note the crucial importance of the fact that some affordances are constructed by the animal itself. In an important, and more complex, subset of such cases, the resulting niche can be seen not merely as a contingent consequence of behavior, but as a kind of quasi-artifact, inasmuch as phenotypic individuals are genetically, morphologically and behaviorally adapted to the construction of specific niches which are integral to the survival and/or reproduction strategy of the species. The term “quasi-artifact” is used here to indicate that, unlike the canonical case of human artifacts, such animal quasi-artifacts need not be produced intentionally in a process guided by representation and conceptualization. Examples of such quasi-artifactual niches are the nests of bower birds, and the dams of beavers. The male bower bird builds and decorates an elaborate nest (bower) to attract females, using attractive objects such as flowers, shells and leaves. The bower forms an integral part of the male’s mating display, and sexual selection by the female is based upon the aesthetic qualities of the bower, as much as upon the behavioral display of the male. Beavers construct, through coordinated and collaborative behavior, dams that serve both as a defence against predators, and as a means to enhance the availability of food. The dams of beavers not only serve as a constructed, quasi-artifactual niche for beavers themselves, but also reproduce the wetland ecology in which many other species thrive. As a final example of the significance of animal quasi-artifacts, we can mention the termite mound, whose material structure is not only integral to the reproductive strategy of this species of social insect, but also constitutes the morphological structure of the colony as a “group organism”. What are the implications of this integration of ecology into evolution for the Neo-Darwinian synthesis? A conservative interpretation would be that the only modification required is that the phenotype, or “vehicle”, be extended to incorporate the quasi-artifactual niche. This is, indeed, the interpretation favored by Dawkins (1982), who employs the terminology of “extended phenotype.” Under this interpretation, the “replicator” remains the gene, and only the gene. In fact, however, it is not only the gene that is copied or replicated. The niche, too, is both reproduced across generations, and serves as a fundamental precondition for genetic replication. The quasi-artifactual niche is thus both a consequence of and an agent in natural and/or sexual selection, and must then be seen as a key ingredient of the evolution of the species-typical genotype. The artifact/niche is not merely part of what is reproduced, but is also fundamental to the process of its reproduction and transmission, since it constitutes a self-made environment for adaptive selection. This co-evolutionary dynamic of reciprocal construction is captured by Laland et al., term “phenogenotypic replication”. It seems, therefore, that the integration of ecological considerations into evolutionary theory, and specifically the existence of animal quasi-artifactual niches, undermines the hard and fast distinction in the modern synthesis between germ-line and soma, genotype and phenotype, “replicator” and “vehicle”. In fact it makes better sense to say that, even granted that the unit of Darwinian selection remains the gene (allele), the “replicator” includes both the artifactual niche, and the nicheadaptive behavioral repertoire of the animal. Such considerations lead us back to Piaget’s more general proposition that behavior is the leading edge and motor of evolution, prompting the conclusion (anticipated above) that the identification by Dawkins of the “replicator” with the unit of selection (the gene, or its hypothesized cultural analogue, the “meme”) is deeply flawed, and that replication can just as well, or even better, be considered as a property of the entire site of selection—including the animal’s behavioral repertoire. In the cases discussed above, the behavioral repertoire of the species includes behaviors that are specifically adapted to the making of the quasi-artifactual niche, and these behaviors in turn support wider repertoires of behavioral strategies exploiting the niche. Quasi-artifactual niches are adaptive precisely because of the behaviors and strategies that they afford—nests are for nesting, and burrows are for burrowing. The quasi-artifactual niche in many cases can be regarded as an extension either of a behavioral repertoire (e.g., male mating display) or of the organism’s morphology (e.g., the bower bird’s bower as functionally equivalent, as an indicator of fitness, to the tail of the peacock). Indeed, we can further ask if it might be fruitful to consider certain species-specific behavioral repertoires, such as birdsong, also to be kinds of animal quasi-artifacts, inasmuch the song of the adults provides a niche within which the singing behavior is epigenetically learned (Marler and Peters, 1982; Sinha, 2004). In general terms, culturally transmitted, specialized behavioral repertoires are not merely biological, but biocultural niches, functionally equivalent to, and constructively integrated with the material dimensions of animal quasi-artifacts. If this argument is accepted, it follows that human natural languages can also be viewed as species-specific, quasi-artifactual, biocultural niches. Pinker (1994), in keeping with his nativist modularist view of the language capacity, emphatically denies that language is an artifact: he regards language as a part of the natural world, and the capacity for language as a part of human nature. We can counter Pinker’s view, however, not by insisting that languages are cultural artifacts tout court, but by regarding language as a biocultural niche and ground for what is unique about human symbolic culture. This conclusion extends the proposals advanced by Odling-Smee and Laland (2009) and Whiten and Erdal (2012), that hominid niche construction created conditions favorable for the emergence and evolution of language, by conceptualizing language itself as a constitutive (indeed the most distinctive) constituent of the evolving human biocultural complex. In other words, the human language capacity, Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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as a psychobiological and organismic property, did not merely evolve within a generalized human socio-cognitive niche, but can be seen as having co-evolved with the linguistic-symbolic niche created by human communication and human praxic action. As I shall argue, this apparently innocuous extension has far-reaching implications for our understanding of the ontology of language and the human language capacity. Language is a species-specific biocultural niche (Sinha, 2009; on which this section is in part based), and the ability to use and acquire it involves not only the replication of phylogenetic adaptation to this niche, but the evolution and replication of an entire symbolically mediated biocultural complex, or what Lotman (1990) called the semiosphere: the ensemble of semiotic resources implicated in the establishment, maintenance and transformation of socio-cultural habitus (Bourdieu 1977). My proposal, in a nutshell, then, is that although other species than humans may properly be said to display behaviors that can be regarded as both cultural and culturally transmitted (Whiten et al., 1999), human culture is distinguished by the predominant place occupied in it by language as a biocultural niche. Language therefore has a dual ontology (Sinha, 2006), as part of biological human species-being, “what it means to be human”, and as the foundational social institution in the Durkheimian sense (Durkheim, 1895). Furthermore, the possession of language imparts this very same dual ontology to our symbolic species (Deacon, 1997; Smolka et al., 1997); perhaps contributing, too, to a distinctively human, semiotic mode of consciousness (Arbib, 2014). Treating language as a biocultural niche yields a new perspective both on the human language capacity and on the evolution of this capacity. First, it unifies, in a non-reductionist fashion, the evolutionary dynamics of human material culture and symbolic culture. As Boivin (2008: 190) has pointed out “Tools, technologies, and other aspects of the material world of humans and their predecessors have largely been seen as the outcome of evolutionary developments, and little attempt has been made to investigate their potential role as selection forces during the course of human evolution” (my emphasis). The same can be said of the biocultural niche of language, which is not separate from the other material and symbolic components or niche-structures that make up the human biocultural complex. The biocultural niche of language is culturally situated, that is, it is dynamically embedded within the entire biocultural complex that includes both symbolic and non-symbolic artifacts. The self-constructed human biocultural complex both favored the emergence and elaboration of language, as proposed by Odling-Smee and Laland (2009: 120); and, because language is co-constitutive of the complex, was fundamentally transformed into a symbolic niche or semiosphere continuous with what we might call the materialartefactual technosphere. We can hypothesize that this process began with proto-language, almost certainly well before the emergence of anatomically and genetically modern humans some 200,000 years ago; and probably in the course of the evolution of Homo erectus from no earlier than 1.8 million years and no later than 500,000 years ago. Recent archaeological discoveries show conclusively that H. erectus individuals had both the motor manipulative skills and the semiotic capacity and motivation to engrave shells at around half a million years ago (Joordens et al., 2014). An obvious question in this context is whether such artifacts can be treated as a proxy for the emergence of evolutionary modern languages; I return to this question in the concluding section. Second, treating language as a biocultural niche means that ecological and epigenetic theories of language learning do not require the organism, as do generative linguistic accounts, to possess an innate Universal Grammar to account for language acquisition. The grammar of the language is not pre-inscribed in a language faculty; rather, grammar is the set of normative conventions regulating action within the semiosphere. This account of language and language learning is thus compatible with usage-based, cognitive-functional theories of language and language acquisition (Tomasello, 1998, 2003). The capacity for constructing a language is, from this perspective, a developing cognitive-behavioral relationship between language user and the constituents of language, just as the capacity for building a nest is a cognitive-behavioral relationship between the builder and the constituents of the nest; and it is this relationship that, in each case, has been selected for in evolution. Human natural languages are dramatically different from the communicative signals and signal systems that are ubiquitous in the animal world. The origin of this difference resides in the fact that only human languages systematically employ symbols rather than signals (Sinha, 2004). The human symbolic capacity emphatically does not take the form of a single design blueprint for grammar. Rather, its productivity and flexibility has eventuated in massive diversity of human languages and human cultures. Rejection of the hypothetical construct of Universal Grammar is consistent with recent comparative linguistic research that highlights dimensions of language variation that have in the past received insufficient attention, and which cast doubt on whether there are any true language universals at all (Evans and Levinson, 2009). Common properties of languages take the form of constraints on variation rather than putative universals, and can be hypothesized to be accounted for by common properties of the human cognitive system, in concert with the transculturally shared communicative intentions of language users. It is true that the capacity to acquire language is a transcultural and species-specific human cognitive capacity, but as I argue below this is an evolutionary outcome consequent on epigenetic plasticity in the extended life course of human infant and child development. 4. Epigenesis, enchrony and evolution Epigenesis and epigenetics are terms referring to inheritance processes and mechanisms, at different levels ranging from the molecular to the organismic, that are controlled or modulated by factors other than those inscribed in the genome (Jablonka and Lamb, 2005). Epigenesis occupies a central position in Piaget’s later work, which was strongly influenced by the theoretical biology of C.H. Waddington (e.g., Waddington, 1953). Piaget considered epigenesis to provide a third way Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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between nativism and environmentalism, and to underpin his constructivist genetic epistemology. Sinha (1988) argued that epigenetic processes provide an integrative bridge between biological and social processes in evolution and development, introducing the notion of epigenetic socio-naturalism. In this section, I outline the integrative relations between niche construction and epigenetic processes in human ontogenesis and its evolution, proposing that this dynamic coupling grounded the emergence of symbolic behaviors. Epigenetic developmental processes in ontogenetic behavioral development are those in which the developmental trajectory and final form of the developing behavior are a consequence as much of the environmental information as of the genetically encoded information. A genetically specified initial behavioral repertoire is subsequently elaborated through experience of a relevant environment, yielding an envelope of potential trajectories and outcomes. The process of elaboration is directional, and once it has taken place the initial plasticity of the embryonic, or unelaborated, repertoire is largely (though not necessarily wholly) lost. In other words, epigenesis involves a developmental transition from relative organismic plasticity and informational openness, to relative rigidity and informational closure. Augmented epigenesis is therefore, advantageous for organisms in which phenogenotypic replication involves a high level of diversity of organism-niche coupling and developmental trajectories. This is particularly relevant in the light of the diversity stressed above in human languages, and the ensuing diversity of human symbolically and linguistically mediated cultures. Regulatory genes augmenting epigenetic openness can therefore be expected to have been phenogenotypically selected for in the evolving human genome, permitting further adaptive selection for domain-specific learning in the biocultural complex, in particular for language learning and for the mimetic learning by observation of both praxic and ritual action (Donald, 1991). The class of organisms with the language capacity (normally developing human beings) can thus be theorized as a phenogenotypic replicator, systemically associated with a wider biocultural complex of interpenetrating semiosphere and technosphere, supported by symbolic and praxic-constructive cognitive capacities. Individual language acquisition and use is situated in the contexts of actuation of these inter-related capacities. What makes human symbolic culture unique is not an innate language acquisition device plus a variety of other species-specific innate cognitive modules, but a generalized semiotic capacity, epigenetically developed from a suite of cognitive capacities largely shared with other species, but attaining higher levels of organization in humans (Deacon, 1997; Piaget, 1945; Sinha, 1988, 2004). This capacity is not inscribed in the human genome, but distributed across the practices and systems co-constituting (with the epigenetically developed human organism) the human phenogenotype. The most fundamental property of these distributed and interlocking systems is, I suggest, enchrony, which is common to practices in both semiosphere and technosphere, and to those frequent cognitive-cultural practices that weave these two spheres together. As yet, we know too little to accept or reject hypotheses regarding the innateness of a specifically syntactic component of the human language capacity. We certainly should not exclude the possibility that the epigenetic processes selected in the evolution of the human biocultural complex include a predisposition for learning syntax; but this does not imply that any such predisposition is or was dedicated from the start exclusively to language. Furthermore, innate adaptations of the perceptionaction system (including the Mirror System: Arbib, 2012) facilitating the epigenetic learning of languages do not imply the innate specification of putative language universals. In an epigenetic perspective, any adaptive developmental predisposition for learning language needs neither to involve direct coding of, nor to be dedicated exclusively to, linguistic structure (Mueller, 1996). Rather, we may hypothesize that epigenetically governed adaptations initially evolved in response to proto-linguistic socio-communicative and symbolic processes, later capturing and re-canalizing behavioral adaptations (such as serial and hierarchical constructive praxis and collaborative joint action) that were initially targeted to other developmental and cognitive domains. The common factor binding these domains together is their enchronic nature, whether involving inter-individual or intraindividual temporal organization and coordination. In the materially productive realm of the technosphere, this involves the evolution of skilled praxic action, and of its learning. In the intersubjective realm of the semiosphere, this involves symbolically mediated skilled symbolic action and interaction (Fusaroli, Gangopadhyay, & Tylén, 2014), and its learning. The learning of these skilled practices is not to be understood as the processing and internalization of an “input” divorced from the learner’s activity. Rather, learning is based in the infant and child’s participation (Goodwin and Goodwin, 2004) in socially organized, enchronically structured practices, supported and constrained by the material and symbolic properties of the co-developing proximal niche. It is to the challenges posed by this socio-culturally organized, diverse, culture- and language-specific developmental process that augmented epigenesis provided the evolutionary answer. 5. Conclusion: a question of timing I conclude by suggesting that augmented epigenesis in modern human ontogenesis was not a necessary condition for the emergence of symbolic capacities and their cognitive prerequisites, which can be traced back at least half a million years. Rather, augmented epigenesis in the time-extended human ontogenetic niche furnished a phenogenotypic mechanism suitable for stabilizing and expanding the diversification and innovation in material and symbolic cultures that first appears in the archaeological record in Southern Africa in the Middle Stone Age, developing between 100,000 and 75,000 years ago (Henshilwood and Dubreuil, 2011; Henshilwood et al., 2011). This diversification and innovation is taken by many archaeologists to be the hallmark of “behavioral modernity”. Criterial for behavioral modernity is the existence of a symbolically mediated culture, in which “individuals understand that artifacts are imbued with meaning and that these meanings are Please cite this article in press as: Sinha, C. Ontogenesis, semiosis and the epigenetic dynamics of biocultural niche construction. Cognitive Development (2015), http://dx.doi.org/10.1016/j.cogdev.2015.09.006
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construed and depend on collectively shared beliefs”; which in turn “explains how human norms and conventions differ from the ritualized behaviors found in non-human primates” (Henshilwood and Dubreuil, 2011: 368–369). As noted above, there is evidence of the engraving by late H. erectus of quasi-geometric designs on shells, around half a million years ago (Joordens et al., 2014). This does not, in itself, warrant the attribution of a full-blown repertoire of behavioral modernity to these hominins. Indeed, such findings highlight the difficulties, both methodological and theoretical, that beset attributions of symbolic mediation of collectively shared beliefs to the cultures of the makers of archaeological artifacts. Nonetheless, these findings compel the conclusion that the time depth of the co-evolution of semiosphere and technosphere is far greater than that of our species. Do they also compel the conclusion that evolutionarily modern, complex language might be equally old? Just such a hypothesis was advanced by Dediu and Levinson (2013), who place the dawn of language at around half a million years ago, with the common ancestor of modern humans and Neanderthals, and suggest that presentday languages have traces of the admixture of Neanderthal languages, just as the genomes of some present-day human populations exhibit traces of interbreeding with Neanderthals1 . There is indeed evidence that Neanderthals manifested some of the practices clustered together as behavioral modernity, including personal ornamentation and funerary practices (d’Errico et al., 2012). However, there is at present no evidence that Neanderthals maintained material-symbolic “techno-complexes”, of the kind characterizing modern human populations in Southern Africa, over extended periods. It is also noteworthy that innovations introduced in the Middle Stone Age, some 80,000 years ago, by Southern African modern humans disappear from the archaeological record at about 60,000 years ago, only to be superseded by new innovations appearing some 15–20,000 years later. Some of these later innovations (from about 44,000 years ago) manifest apparent continuity with historic hunter-gatherer societies (d’Errico et al., 2012). It seems, then, that the human cultural “ratchet effect” (Tomasello, 1999), by means of which cultural innovations are preserved, transmitted and serve as the foundation for subsequent innovation, may not have been fully in place until quite late in the evolution of our species. The earliest evidence of an extended human childhood with a life history profile similar to that of contemporary children, based on analysis of the teeth of an early Homo sapiens child, is from 160,000 years ago (Smith et al., 2007). If we accept with Smith, Toussaint, Reid, Olejniczac, and Hublin (2007) that this life course was not shared by Neanderthals, it is a plausible inference that the augmented epigenetic developmental plasticity of our species was also not shared in the same degree by any other (extinct) hominin species, but evolved either as part of, or subsequent to, modern human speciation. What specific advantage, then, might species-specific augmented epigenesis have conferred, and with what consequences? Perhaps the decisive selective advantage conferred by augmented epigenesis had not only to do with the faithful copying and transmission of specific material and symbolic practices, important as this may be; but also with the role of learning and teaching in generating and tracking cycles of expansion and stabilization of the symbolic space of social and cultural structure, brought into being by the evolution of the all-pervasive, linguistically grounded semiosphere. 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1 The shell engravings described by Joordens et al. were found in Java; whereas Dediu and Levinson theorize about the language capacities of an African common ancestor of modern humans and Neanderthals.
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